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Abstract: Carbon storage by trees and soil in urban areas is of increasing interest as a potential greenhouse
gas mitigation measure. Our objectives were to (1) quantify carbon accumulation in above- and below-ground
tree biomass, organic layer and mineral soil (0–5 cm) of a 27-year-old planted forest in Auckland and (2)
compare the sequestration potential of urban trees with natural shrublands and forests in New Zealand. A
mixed-species allometric equation for urban-grown native trees was developed on the basis of the tree biomass
of 21 trees belonging to four species (Corynocarpus laevigatus, Kunzea ericoides, Pittosporum eugenioides,
P. tenuifolium). Our allometric equation and a recently developed mixed-species equation for New Zealand
native forest species produced similar results. A total of 45.9 Mg C ha–1 was stored in above- and below-ground
tree biomass in our Auckland park, equating to an average annual carbon sequestration rate of 1.7 Mg C ha–1.
This rate is within the range reported for New Zealand shrublands. Despite the adverse conditions posed by
some urban environments (e.g. poor-quality soil, contamination), planted urban forests can sequester carbon
at similar rates to natural vegetation in New Zealand.
Keywords: allometric equations; carbon sequestration; planted forest; root biomass; tissue and wood chemistry
clearance of the urban forest and at some time previous to per tree species). The organic layer was sampled within 20
our involvement; by the time we reached the site much of it × 25 cm quadrats and mineral soil samples were taken with
had already been cleared. We were given a brief opportunity an auger from 0–5 cm depth. Mineral soil samples (only one
to visit the site and identify trees we would like to analyse. sample per tree species) were also taken, at 50–55 cm depth,
The trees were extracted and stockpiled to the contractors’ to estimate the amount of carbon stored at the time of tree
timetable and we were then given 3 days on which we could planting. Soil bulk density was calculated as the quotient
work on them. No bulk storage facilities were available to us, of soil dry weight (105°C, 48 h) and soil volume. The stone
so all the basic measurement and sample collection was carried content of the infill material was less than 5%.
out on site. This is also why it was not feasible to separate the The biomass of the organic layer was estimated by
foliage from the branches. oven-drying the material at 105°C for 48 h after removing
The vegetation survey consisted of four transects parallel a homogeneous subsample for carbon analysis. Mineral soil
to each other, from the top of the slope to the bottom. Transect samples and the forest-floor subsamples were oven-dried at
lengths varied from 35 to 50 m, depending upon the width 40°C and ground (ZM 200, Retsch GmbH, Haan, Germany).
of the vegetation at that point. All trees within 1 m of either Mineral soil samples were passed through a 2-mm sieve
side of the transect line were identified and their diameter at before grinding.
1.4 m above ground measured. A general indication of tree
height was recorded. Wood density and carbon analysis
The discs, sections of roots, and crown samples were weighed,
Tree selection air-dried, and reweighed. The volume of the disc and root
As part of the project, the contractors agreed to dig up, in sections was determined by water displacement. The samples
their entirety, 21 trees of four native species to measure were then dried at 105°C until constant weight was achieved
the above- and below-ground biomass accumulation. The (typically 4 days) to estimate both fresh density and dry wood
chosen species – karaka (Corynocarpus laevigatus) (n = 6), density.
kānuka (Kunzea ericoides) (n = 5), lemonwood (Pittosporum Plant and soil material were analysed for carbon
eugenioides) (n = 6), and kōhūhū (P. tenuifolium) (n = 4) – are concentration by dry combustion using an elemental analyser
all widely planted in the Auckland Region in revegetation and (True Spec, LECO Corporation, St. Joseph, Michigan, USA).
restoration projects (Wilcox 2012). Kunzea ericoides can grow Every 10th sample was analysed twice. The analytical precision
into a canopy tree (up to 20 m) whereas the other species are was 1.3% for plant and 0.15% for soil material.
smaller trees (up to 15 m) (Wardle 1991). Proc GLM (SAS version 9.2, SAS Institute Inc., Cary,
Due to access problems we were only able to select trees NC, USA) was used to identify differences in tree and tissue
from certain areas. As far as was possible, individuals typical characteristics among species.
of the height and growth form for that species were selected.
Thus, we ignored obviously suppressed individuals and trees Tree and stand carbon accumulation and sequestration
on the edge that often had a different growth form to those Tree felling is often not an option to derive tree carbon
in the interior. stocks, particularly in urban areas. However, in this case
we were provided with mostly complete trees that had been
Tree measurements and sample preparation mechanically extracted from the ground. The amount of carbon
The trees were measured for their height as well as diameter stored in each tree component was estimated by multiplying
at breast height (1.4 m; DBH) above what had been ground carbon concentrations and the mass (dry weight) of a given
level. The root plate was cut off, and attached soil and stones component. The total tree carbon is the sum of carbon stored
carefully removed. In the extraction process, it was inevitable in above- and below-ground tree components. The proportion
that fine and medium roots were lost. However, from our of below-ground carbon to total tree carbon was calculated as
observations, the majority of the major roots were intact; usually the quotient of root carbon and total carbon stocks.
it was the distal ends that were missing. The stem was cut up From these data we were also able to develop an allometric
into manageable sections and the crown (including branches equation that could be applied to the rest of the trees in
and leaves) separated; the crown was defined as the point at Newmarket Park. We also had other survey data on planted
which multi-branching of leaf-bearing branches occurred. forests in the Auckland Region, many planted during the same
Disks were cut from the stem of each tree for more detailed time period. From these data, species-specific growth curves
laboratory analysis. Each part of the tree was weighed to an were constructed to estimate individual tree heights for the
accuracy of 0.1 kg. measured trees.
Each section of each tree was then chipped using a The amount of carbon stored in the organic layer was
commercial tree chipper. Approximate 5-kg samples of calculated as the product of carbon concentration and the dry
wood chips from the roots, stem and crown of each tree were weight of the organic layer. Mineral soil carbon density (Mg C
retained for laboratory analysis. The approach was to start the ha–1) was estimated by multiplying soil carbon concentration
chipping, allow chips to begin to be produced, and then collect with bulk density and soil depth. The amount of carbon stored
the subsequent 5 kg. This was to avoid any contamination at 50–55 cm depth (carbon density at the time of tree planting
between samples. The wood chip material was subsampled, = baseline carbon density) was subtracted from the carbon
freeze-dried, and then ground to a fine powder using a ultra density at 0–5 cm depth to calculate the accumulation of soil
centrifugal mill (ZM 200, Retsch GmbH, Haan, Germany). carbon since planting.
Table 1. Summary of comparisons among species for the tree and soil characteristics at Newmarket Park, Auckland, New
Zealand. Values are means ± standard deviation. Significance * P < 0.05; ** P < 0.01; *** P < 0.001; ns – no significant
difference.
__________________________________________________________________________________________________________________________________________________________________
Tree height (m) 6.7 ± 0.5 11.3 ± 0.4 10.5 ± 0.6 9.2 ± 0.4 ***
Tree diameter (cm) 12.0 ± 0.8 12.3 ± 2.3 15.0 ± 0.6 13.6 ± 1.3 *
Tree biomass (kg) 46.2 ± 10.5 89.6 ± 15.3 166.1 ± 25.5 113.6 ± 39.4 **
(dry weight)
Proportion of total biomass
Crown (%) 30.3 ± 2.9 15.8 ± 1.1 20.1 ± 1.4 22.5 ± 2.6 **
Stemwood (%) 51.1 ± 2.6 71.7 ± 1.6 60.1 ± 2.8 62.5 ± 1.9 ***
Roots (%) 18.6 ± 3.0 12.5 ± 1.4 19.7 ± 1.7 15.0 ± 2.3 ns
Root-to-shoot ratio 0.23 ± 0.04 0.19 ± 0.05 0.25 ± 0.09 0.29 ± 0.08 **
Wood density
Stemwood (g cm–3) 0.47 ± 0.01 0.68 ± 0.01 0.63 ± 0.00 0.66 ± 0.01 ***
Roots (g cm–3) 0.50 ± 0.01 0.70 ± 0.01 0.65 ± 0.04 0.71 ± 0.07 **
Carbon concentration
Crown (%) 44.9 ± 1.4 49.6 ± 1.5 46.6 ± 0.4 47.6 ± 0.6 ***
Stemwood (%) 45.2 ± 0.1 47.0 ± 0.3 45.9 ± 0.2 45.6 ± 0.1 ***
Root (%) 43.4 ± 2.4 46.3 ± 1.2 43.8 ± 1.7 44.7 ± 0.5 ns
Tree carbon content (kg tree–1) 15.1 ± 3.2 33.5 ± 5.8 62.1 ± 9.5 42.9 ± 15.0 **
Proportion of total tree carbon
Stemwood + crown (%) 76.7 ± 4.0 84.5 ± 1.7 77.6 ± 1.9 82.2 ± 2.8 **
Roots (%) 23.4 ± 4.0 15.5 ± 1.7 22.4 ± 1.9 17.8 ± 2.8 ***
Organic layer1
Biomass (dry weight) (Mg ha–1) 16.6 ± 3.1 22.5 ± 4.1 15.8 ± 3.5 16.2 ± 4.2 ns
Carbon concentration (%) 37.4 ± 1.6 40.5 ± 0.7 34.2 ± 5.5 33.5 ± 1.5 **
Carbon density (Mg C ha–1) 6.2 ± 0.8 9.1 ± 1.3 5.4 ± 1.6 5.5 ± 1.1 ***
Mineral soil1
0–5 cm
Carbon concentration (%) 5.8 ± 0.4 5.7 ± 0.7 5.3 ± 0.5 7.7 ± 0.4 ***
Carbon density (Mg C ha–1) 25.5 ± 1.7 25.2 ± 3.2 23.3 ± 2.0 32.3 ± 1.5 ***
50–55 cm
Carbon concentration (%) 0.5 0.8 0.8 1.9 ***
Carbon density (Mg C ha–1) 2.9 5.3 5.3 12.9 ***
__________________________________________________________________________________________________________________________________________________________________
1
n = 3 per tree species
216 New Zealand Journal of Ecology, Vol. 38, No. 2, 2014
Tree carbon content (kg tree–1) = −2533.5 volume3 + 1323.2 values for asp based on those presented in Beets et al. (2012).
volume2 + 117.59 volume, where volume (m3) = tree height There is clearly much variability between species for asp, with
× tree basal area. no obvious patterns. However, hardwood and softwood species
tend to have different wood characteristics. In our study we only
This polynomial function was then used to estimate the
had hardwood species thus, for the branches and foliage terms,
carbon storage at the stand level using the transect data.
we used the mean asp based on the hardwood species listed in
Beets et al. (2012). We identified that variation in the asp value
The second allometric equation developed for Newmarket
of the ‘stem and branch carbon’ term has the greatest effect on
Park is based on a power function. This equation has three
overall tree carbon estimation; this also corresponded to our
components (stem and branches, crown and roots) and
‘trunk’ measurement. For each of our species, we developed,
combined these components such that we could estimate
through successive approximation, a ‘stem and branch carbon’
either the above-ground carbon or the total tree carbon. The
asp value, which was used to estimate the carbon content of
equation took the form:
each individual of a species.
Tree carbon content (kg C) = 0.0023DBH3.3885 +
Tree carbon content (kg C) = asp (DBH2 × Height)0.936 +
0.0121DBH2.5276 + 0.009DBH2.4966.
0.0197 DBH0.936 + 0.0148 DBH1.595.
This corresponds to: stem and branches + crown + root.
The asp was modified until the sum of the carbon content
for individuals of that species was equivalent to the measured
Both the polynomial (Proc GLM) and power function (Proc
above-ground value (see Table 2 for the asp values).
POWER) were developed using SAS version 9.2 (SAS Institute
Inc., Cary, NC, USA). The variables were tree volume and
Stand carbon accumulation and sequestration rates
tree diameter, respectively.
The vegetation survey included 12 species that had been
Comparison with the allometric equations of Beets et al. planted. Kunzea ericoides was present in the replanted area but
(2012) not along the four transects. Tree density varied considerably
among species, reflecting differing survival rates or initial
Beets et al. (2012) developed allometric equations (using power
densities (Table 3). Over the 27 years of its existence this forest
functions) for New Zealand forest tree species on the basis
stand accumulated a total 45.9 Mg C ha–1 in above- and below-
of biomass data of approximately 140 trees. The authors also
ground tree biomass (Table 3). The tree carbon sequestration
demonstrate that, for improved precision, parameterisation
rate was 1.7 Mg C ha–1 year–1, with considerable differences
of the species-specific equations varies between species
among species (Table 3). Including the amount of carbon
(Beets et al. 2012). Power functions are the most widely used
stored in the organic layer (6.5 Mg C ha–1) and mineral soil
functions for allometric equations (Picard et al. 2012), but in
(0–5 cm; 20.2 Mg C ha–1 corrected for mineral soil carbon
our analysis we found that a polynomial function provided the
density at the time of tree planting), a total of 72.7 Mg C ha–1
best estimation of total tree carbon content. It is thus important
was stored in this 27-year-old urban park (Table 3).
to compare our results with those that used allometric equations
The ‘Newmarket power function’ and ‘Beets general
based on power functions.
equation’ were also applied to the Newmarket Park vegetation
Presented in Table 2 are results for both above-ground and
survey data to estimate overall carbon accumulation at the
total carbon based on our (destructive) analysis together with
site (Table 4).
a set of additional analyses. The ‘Beets general mixed-species
equation’ (Beets et al. 2012) is expressed as:
Tree carbon content (kg C) = 0.0162 (DBH2 × Height)0.943 Discussion
+ 0.0175 DBH2.2 + 0.0171 DBH1.75.
Carbon concentrations of tree tissues and carbon
The ‘Beets modified equation’ is based on the above equation accumulation on a per-tree basis
but using a species-specific ‘a’ parameter (asp). In our study The higher carbon concentration in crown tissue is explained
we did not separate foliage from branches thus we retained by the high proportion of foliage. Leaves and needles have
Table 2. Comparison of the measured carbon content with the estimated carbon content derived from allometric equations.
a sp = species-specific ‘a’ parameter.
__________________________________________________________________________________________________________________________________________________________________
Table 3. Stem density, basal area, carbon accumulation and sequestration of trees, organic layer and mineral soil carbon density (0–5 cm).
Stem density and basal area result from the vegetation survey. The tree carbon values were estimated using the ‘polynomial function’
derived from biomass data of 21 trees felled in Newmarket Park, Auckland, New Zealand.
__________________________________________________________________________________________________________________________________________________________________
1
Species Stem density Basal area Carbon accumulation Carbon sequestration
(ha–1) (m2 ha–1) (Mg C ha–1) (kg C ha–1 year–1)
__________________________________________________________________________________________________________________________________________________________________
Table 4. Site estimation of carbon content at Newmarket Park, Auckland, New Zealand. Values derived from three allometric
equations are provided for both individual species and overall estimate of carbon.
__________________________________________________________________________________________________________________________________________________________________
been reported to have higher carbon concentrations compared data on tree tissue carbon.
with stem wood and roots (e.g. Bert & Danjon 2006; Mello The amount of carbon accumulated in individual trees
et al. 2012). Stem-wood carbon concentrations measured in differed considerably among tree species (Table 1). Small
the four New Zealand native tree species (45.2–47.0%) were tree size and low wood density explain the low amount of
at the lower end of values reported for angiosperms from carbon stored in Corynocarpus laevigatus. Total-tree carbon
across the globe (47.7 ± 0.3%) (Thomas & Martin 2012). In accumulation is also influenced by species-specific allocation
most New Zealand studies a constant conversion factor of strategies (e.g. Campioli et al. 2010). A higher proportion of
0.5 is used to convert dry biomass into carbon stocks (e.g. stem and root biomass resulted in higher tree carbon stocks in
Beets et al. 2012; Carswell et al. 2012). Using the general Kunzea ericoides, Pittosporum eugenioides and P. tenuifolium.
default value of 0.5, the carbon accumulation of individual The allocation into slowly decomposable carbon pools such
trees in Newmarket Park would have been overestimated by as stems and roots also implies that the carbon sequestration
3.0–4.8%. This illustrates the need to obtain species-specific capacity of these species in the long term is higher than the
218 New Zealand Journal of Ecology, Vol. 38, No. 2, 2014
carbon sequestration capacity of Corynocarpus laevigatus. regenerating Kunzea ericoides stand of similar age was 17 330
Roots as a percentage of total tree biomass and the root-to- stems ha–1 (Carswell et al. 2012). Growth rates might also have
shoot ratio in our study were in general lower than the values been reduced as trees were planted on soils and sediments
reported by Phillips and Watson (1994) and Hart et al. (2003). characterised by very low carbon and nitrogen contents. Despite
For example, live fine and coarse roots were 22% of total live these adverse conditions this planted urban forest sequestered
hard beech (Nothofagus truncata, 6–80 cm in diameter) biomass carbon at a similar rate to natural vegetation in New Zealand.
with a root-to-shoot ratio of 0.28 (Hart et al. 2003). The total The amount of carbon stored in the organic layer (5–9
root biomass in six 14-year-old Pinus radiata trees (21–31 cm Mg C ha–1) at Newmarket Park was slightly higher than
in diameter) represented 30% of the total tree biomass and had values measured across 10 urban forests sites (2–6 Mg C
a root-to-shoot ratio of 0.43 (Phillips & Watson 1994). Kunzea ha–1) in Auckland (Wang 2012). Higher organic (litter)
ericoides (32 years old, 12.7 cm in diameter) growing in the layer carbon stocks (8.9 ± 2.2 Mg C ha–1) were found in the
East Coast hills had a root biomass (roots > 2 mm) of 20.1 Leptospermum scoparium – Coprosma propinqua dominated
kg tree–1 (Watson et al. 1995) compared with 15.4 kg tree–1at shrublands investigated by Coomes et al. (2002). For two
Newmarket Park. Owing to the lack of species-specific root species, Corynocarpus laevigatus and Kunzea ericoides,
biomass data, a root-to-shoot ratio of 0.25, based on the IPPC the organic layer- and mineral-soil-associated carbon pools
guidelines for mature temperate broadleaved forests, is often exceeded the amount of carbon stored in tree biomass. Carbon
used to account for below-ground biomass in New Zealand stored in soil can exceed above-ground carbon and the
trees (e.g. Beets et al. 2012). importance of soil carbon storage has recently been highlighted
Some of the differences in root biomass can be explained in a study on carbon stock in urban ecosystems (Edmondson
by variations in root extraction methods or data collection et al. 2012). Our results suggest that the amount of soil organic
methods. Our lower root biomass and root-to-shoot ratio may carbon is tree species dependent. The effect of tree species on
partly be explained by an underestimation of fine roots due to carbon accumulation in soils is explained by the quality and
the root extraction procedure. However, trees grown in urban amount of litterfall and root residues (e.g. Finzi et al. 1998;
environments may allocate less biomass and carbon to roots. A Schulp et al. 2008). The fact that Kunzea ericoides, which had
recent study showed that urban-grown Quercus rubra seedlings the highest carbon stock in the organic layer, did not have the
had a lower root-to-shoot ratio than seedlings grown in remote highest tree biomass stocks implies that there are differences
areas (Searle et al. 2012). The decrease in biomass allocation in litter decomposition rates between species.
to roots has been explained by higher growth temperatures in
urban areas (Searle et al. 2012). Chen et al. (2013) associated Comparison of allometric equations
lower fine root biomass in urban Pinus massoniana forests The polynomial function worked well when applied over the
with higher N depositions in urbanised areas. whole Newmarket Park dataset, despite the varying timber
characteristics. However, for individual trees the accuracy
Comparison with carbon sequestration rates elsewhere varied considerably. Our polynomial function requires only
in New Zealand diameter and height. As these tree characteristics are easy
Carbon accumulated in above- and below-ground tree biomass to measure, tree carbon storage across a wider scale could
at Newmarket Park (45.9 Mg C ha–1) was within the range be evaluated. Still, this allometric equation has to be further
reported for New Zealand shrubland studies. For example, improved by measuring additional tree species and sites. Using
Coomes et al. (2002) estimated 48.6 ± 13.5 Mg C ha–1 (above- this allometric equation for trees outside the diameter range of
and below-ground biomass) in Leptospermum scoparium – trees investigated in this study may not provide reliable carbon
Coprosma propinqua dominated shrublands growing north values, as polynomial functions are subject to large errors when
of Christchurch. An above-ground biomass of 34 Mg C ha–1 extrapolated outside the measured range (Picard et al. 2012).
was reported for different types of New Zealand shrubland The comparison between our measured tree carbon
(Beets et al. 2009 cited in Carswell et al. 2012). Between 12 content and other allometric equations revealed that the more
and 31 Mg C ha–1 were estimated in overseas urban forests generalised equations did less well at estimating tree carbon
under similar climatic conditions to Newmarket Park (Davies content than did the more ‘optimised’ equations. The ‘Beets
et al. 2011; Hutyra et al. 2011; Strohbach & Haase 2012). general mixed-species equation’ with an added 25% for root
The sequestration rate of 1.7 Mg C ha–1 year–1 determined carbon provided a good approximation of the measured total-
for Newmarket Park was within the range of values reported tree carbon values. This suggests that where both height and
for New Zealand shrublands. The rate of carbon sequestration diameter measurements are available, this equation would
for young (age < 50 years) Kunzea ericoides and coastal be a good approach for non-destructively estimating total
broadleaved stands was 2.3 Mg C ha–1 year–1 (Carswell et al. carbon content of a New Zealand mixed forest. Surprisingly
2012). A carbon sequestration rate of 2.1 ± 0.2 Mg C ha–1 this allometric equation did least well at estimating the above-
year –1 for approximately 40-year-old Leptospermum ground carbon, underestimating by some 11%. If height data
scoparium – Kunzea ericoides stands across six sites was are not available, then an allometric equation based on power
reported by Trotter et al. (2005). In contrast, the sequestration functions and diameter may well prove to be reasonably robust
rate of a Pinus radiata plantation forest over a rotation cycle is as shown for North American tree species (Jenkins et al. 2004).
about 8 Mg C ha–1year–1 (Maclaren 2000). Our sequestration The other feature to emerge from these results is that, except
rate was lower than the average sequestration rate of 2.8 C where the equations are species-specific, these equations are
ha–1year–1 by urban trees in the United States (Nowak et al. quite imprecise at estimating the carbon content of individual
2013). trees at a site. However, when applied to the overall site
The slightly lower sequestration rates for the planted estimation, the applicable equations all provided a similar range
forest at Newmarket Park can partly be explained by the of values (Table 4). It might be expected that the ‘polynomial’
comparatively low tree density. Only 2212 stems ha–1 were and ‘power’ functions would provide a similar outcome as
measured in the stand. In contrast, tree density in a naturally they are based on local data. Interestingly, the ‘Beets general
Schwendenmann, Mitchell: Carbon accumulation in an urban park 219
equation’ produced a similar result, suggesting the wide in European cities. Biology Letters 5: 352–355.
applicability of this equation. If many more species can be Gaffin SR, Rosenzweig C, Kong AYY 2012. [Correspondence]
parameterised then, clearly, using species-specific allometric Adapting to climate change through urban green
equations will be the optimal solution. infrastructure. Nature Climate Change 2: 704.
Hart PBS, Clinton PW, Allen RB, Nordmeyer AH, Evans G
2003. Biomass and macronutrients (above- and below-
Acknowledgements ground) in a New Zealand beech (Nothofagus) forest
ecosystem: implications for carbon storage and sustainable
The former Auckland City Council provided a small grant forest management. Forest Ecology and Management
towards this project and facilitated the provision of trees; funds 174: 281–294.
from The University of Auckland School of Environment PBRF Hutyra LR, Yoon B, Alberti M 2011. Terrestrial carbon stocks
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the unnamed digger drivers who carefully extracted the trees Comprehensive database of diameter-based biomass
for us and placed them in a neat stockpile; Peter Crossley for regressions for North American tree species. General
chainsawing the trees with great skill and efficiency; Gabriella Technical Report NE-319. Newton Square, PA, USDA,
Ezeta-Ramos for on-site assistance; Sara Howarth of Tonkin Forest Service, Northeastern Research Station. 45 p.
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Received 3 May 2013; accepted 16 November 2013
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